Turtles (Reptilia: Testudines) Of The Ardis Local Fauna Late Pleistocene (Rancholabrean) Of South Carolina Curtis C Bentley and James L. Knight South Carolina State Museum, 301 Gervais Street, P.O. Box 100107 Columbia, South Carolina 29202-3107 ABSTRACT- The Ardis local fauna (late Pleistocene) was collected from a group of interconnecting sediment-filled solution cavities, located in the Giant Cement Quarry near Harleyville, Dorchester County, South Carolina. Fossil material from the lowermost levels and the extreme upper layer of the deposit have been radiocarbon dated at 18,940 ± 760 and 18,530 ± 725 y.b.p., respectively. These dates are considered con- temporaneous within present resolution. Approximately ninety verte- brate taxa were collected from the site. Fourteen were species of turtles, including eight not previously reported from the Pleistocene of South Carolina. Among these is the southeasternmost occurrence of Emy- doidea blandingii. This record, in conjunction with other vertebrate fos- sils from the site, suggests a north-south dispersal route of species along the Atlantic Coastal Plain during interglacial-glacial transitions. Geo- graphically isolated eastern and western populations of Emydoidea blandingii may have existed during the maximum advance of the Lau- rentide ice sheet. Unusually complete fossils of large box turtles recov- ered from the site corroborate the previously suggested synonymy of the extinct Terrapene Carolina putnami with T c. major. The fossil turtle community of the Ardis local fauna has no modern analogue. Like the Ardis mammals, it comprises a "disharmonious" fauna which suggests that, during the height of the Wisconsinan glaciation, the region experi- enced a more equable climate than that of today . The Ardis local fauna has yielded approximately 90 species of late Pleistocene fossil vertebrates from the Coastal Plain of South Carolina, includ- ing a substantial mammalian (Bentley et al. 1994), avian, reptilian, amphibian, and fish faunas currently under study. We present data on the Ardis turtle col- lection (Appendix I), the largest Rancholabrean fauna reported from the state. Dobie and Jackson (1979) and Roth and Laerm (1980) reported fossils of late Pleistocene age from Edisto Island, the only other Pleistocene fossil turtle fauna from South Carolina described to date. Among the Edisto Island fauna were ten Curtis C. Bentley and James L. Knight taxa of turtles, including Gopherus sp. and Malaclemys terrapin which were not recovered from the Ardis local fauna and possibly three species of Pseudemys, P. floridana and/or P. concinna, and P nelsoni. The Ardis local fauna was discovered in a large open-pit mine, operat- ed by the Giant Cement Company, located 5 km NNE of Harley ville, Dorchester County, South Carolina (33° 14'N, 80° 26'W). Quarry operations exposed San- tee Limestone (middle Eocene) and the clay-rich Harleyville Formation (late Eocene) which underlie Plio-Pleistocene surficial deposits (Ward et al. 1979, Harris and Zullo 1991). Locally, groundwater differentially dissolved the Santee Limestone in its upper portions, so that many solution cavities contacted and penetrated the overlying Harleyville Formation and thereby opened several of the cavities to the Pleistocene surface (Bentley et al, 1994). The radiocarbon dates of the Ardis material place the time of deposition at or near the height of the Wisconsinan glaciation (Bowen 1988, Tushingham and Peltier 1993). Fur- ther discussion on the geology, dating methods of the Ardis fossil material, pre- vious fossil collections from the quarry, fossil collection procedures, and a local- ity map are available in prior publications (Bentley and Knight 1993, Bentley et al. 1994). TAPHONOMY At least part of the fossil assemblage collected from inside the solution cavities at the Ardis site appears to represent an obrution deposit, the very rapid burial of intact organisms (Brett 1990) in which many of the specimens exhibit incipient decay. Surface openings leading to the cavities varied from a gentle downward slope to a vertical shaft, generally allowing the Pleistocene fauna ease of ingress and egress. This permitted animals to enter the cavities in three dif- ferent ways: (1) "walk-in" taxa, which may have used the site for estivation/hibernation or as denning sites and hunting grounds, for example muskrats, mink, and woodrats (Bentley et al. 1994); (2) "wash-in" taxa from the surface, either alive or dead, which applies most readily to large animals known only from isolated remains e.g., Mammut sp., Bison sp., Equus sp. (Bentley et al. 1994), that would have been unable to enter the cavities during life; and (3) "fall- in" taxa which fell into exposed verticle shafts, fossil accumulations resulting from this type of natural trap are well documented (e.g., Webb 1974). Because of the interconnecting "tunnel-like" nature of the cavities, a single episodic event could produce differing water velocities within the cavities and different rates of deposition. Seasonal flooding, depending on the intensity, may have simultaneously smothered living animals within the cavities and buried or reworked those that had died just prior to, or in a preceding, deposi- tional event. Consequently, specimens incompletely or shallowly buried during an event with a low sedimentation rate (low energy) could be completely or par- tially exhumed and reburied by a succeeding event. This resulted in the preser- Turtles vation of specimens in various orientations to the bedding planes (Fig. 1), vari- ous degrees of disarticulation, and the occasional mixing of individual elements. An articulated Emydoidea blandingii specimen preserved in its life position with axial skeletal elements inside, indicates little or no decay prior to its final burial (Fig. 2). This preservation suggests the turtle was buried quickly in a high ener- gy, high sedimentation environment (Brett and Speyer 1990), resulting in burial deep enough to avoid reworking during subsequent episodic events. Several articulated turtles were collected with limbs and skulls preserved within the shells in various orientations to the bedding plane. A high energy hydrological environment before or shortly after death would likely explain the various orien- tations observed in well preserved specimens. Retention and preservation of limb elements, cervical and caudal vertebrae, and skulls inside the shell may reflect a withdrawal by the turtles in response to a catastrophic event. Fig. 1 Emydoidea blandingii, only the carapace (.547) was preserved, ventral side up (side view), among clay clasts from the surrounding Harleyville Forma- tion. This illustrates the hydrodynamic effect upon some specimens prior to final burial. Curtis C. Bentley and James L. Knight Fig. 2 Complete E. blandingii (.546) in situ, with axial skeleton preserved inside the shell, indicating a "withdrawal response" and final burial prior to any signif- icant decay. MATERIALS AND METHODS Morphological terminology used in this paper is taken from Carr (1952), Ernst and Barbour (1989), Holman (1967, 1977, 1985), Holman and Grady (1987), and Preston (1979). Taxonomy follows Conant and Collins (1991). Morphological comparisons of Recent skeletons to fossil material were made against available specimens in the Florida Museum of Natural History and the South Carolina State Museum collections. Additionally, specimens from The University of Michigan Museum of Zoology of Emydoidea blandingii, UMMZ 155047-155054, Clemmys guttata, UMMZ 51235, 51236, 51240-51242, 159219, 155001, 155002, and Clemmys muhlenbergii, UMMZ 77140 and, 130840 were studied. Most of the specimens in the South Carolina State Museum collections are deposited under the base number of S.C. 94.10. and for brevity, are refer- enced in the text only by the digits following this base number. Specimens accessioned separately are designated by the institutional prefix of SCSM. Fos- sil specimens deposited in the National Museum of Natural History and the Florida Museum of Natural History are designated by USNM and UF, respec- tively. Turtles SYSTEMATIC PALEONTOLOGY Testudines Kinosternidae Material: 1 right xiphiplastron (.25); 16 peripherals (.26-.44); 2 humeri (.45-. 46); 2 partial jaw rami (.755-. 756). Remarks: These fossil elements could only be identified with confidence to fam- ily. Kinosternon subrubrum - Eastern Mud turtle (Lacepede, 1788) Material: 3 nuchals (.11 -.13); 2 right, 3 left hyoplastra (.6-. 10); 5 left hypoplas- ia (.1-.5). Characters used for identification: The hyoplastron of Kinosternon subrubrum can be separated from other North American Kinosternon and Sternotherus because the axillary notch is narrower, and from Kinosternon baurii because the axillary notch is wider and shallower (Holman 1985). K. subrubrum hyo- and hypoplasia differ from Sternotherus odoratus in that the elements are shorter lat- erally than medially in S. odoratus (Preston 1979). Characters used to identify nuchal material are discussed by Holman (1975). In addition, nuchals of K. sub- rubrum can be distinguished from nuchals of S. odoratus because the anterior lip of the nuchal, viewed anteriorly, is nearly straight in K. subrubrum. Nuchals of S. odoratus, viewed anteriorly, have a decided arc. Remarks: The eastern mud turtle inhabits a variety of shallow slow to non-mov- ing bodies of water with a soft substrate, such as swamps, ponds, marshes, wet meadows, and lagoons (Ernst and Barbour 1989). Kinosternon subrubrum today ranges from southern Massachusetts and Pennsylvania